Journal of Plant Physiology 167 (2010) 1052–1060
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Journal of Plant Physiology
journal homepage: www.elsevier.de/jplph
Photosynthesis and photoprotection in coffee leaves is affected by nitrogen and
light availabilities in winter conditions
Marcelo F. Pompelli
1
, Samuel C.V. Martins, Werner C. Antunes, Agnaldo R.M. Chaves, Fábio M. DaMatta
∗
Departamento de Biologia Vegetal, Universidade Federal de Vic ¸ osa, 36570-000 Vic ¸ osa, MG, Brazil
article info
Article history:
Received 15 December 2009
Received in revised form 4 March 2010
Accepted 4 March 2010
Keywords:
Antioxidant enzymes
Coffea
Oxidative stress
Photoinhibition
Xanthophyll cycle
abstract
Coffee is native to shady environments but often grows better and produces higher yields without shade,
though at the expense of high fertilization inputs, particularly nitrogen (N). Potted plants were grown
under full sunlight and shade (50%) conditions and were fertilized with nutrient solutions containing
either 0 or 23 mM N. Measurements were made in southeastern Brazil during winter conditions, when
relatively low night temperatures and high diurnal insolation are common. Overall, the net carbon assim-
ilation rate was quite low, which was associated with diffusive, rather than biochemical, constraints. N
deficiency led to decreases in the concentrations of chlorophylls (Chl) and total carotenoids as well as
in the Chl/N ratio. These conditions also led to qualitative changes in the carotenoid composition, e.g.,
increased antheraxanthin (A) and zeaxanthin (Z) pools on a Chl basis, particularly at high light, which was
linked to increased thermal dissipation of absorbed light. The variable-to-maximum fluorescence ratio
at predawn decreased with increasing A + Z pools and decreased linearly with decreasing N. We showed
that this ratio was inadequate for assessing photoinhibition under N limitation. Expressed per unit mass,
the activities of superoxide dismutase and glutathione reductase were not altered with the treatments.
In contrast, ascorbate peroxidase activity was lower in low N plants, particularly under shade, whereas
catalase activity was lower in shaded plants than in sun-grown plants, regardless of the N level. Glutamine
synthetase activity was greater in sun-grown plants than in shaded individuals at a given N level and
decreased with decreasing N application. Our results suggest that the photoprotective and antioxidant
capacity per amount of photons absorbed was up-regulated by a low N supply; nevertheless, this capac-
ity, regardless of the light conditions, was not enough to prevent oxidative damage, as judged from the
increases in the H
2
O
2
and malondialdehyde concentrations and electrolyte leakage. We demonstrated
that N fertilization could adequately protect the coffee plants against photodamage independently of the
anticipated positive effects of N on the photosynthetic capacity.
© 2010 Elsevier GmbH. All rights reserved.
Abbreviations: A, antheraxanthin; APX, ascorbate peroxidase; Car, carotenoid;
CAT, catalase; Chl, chlorophyll; C
i
/Ca, internal-to-ambient CO2 concentration ratio;
D, fraction of the absorbed PAR dissipated as heat; DEPS, de-epoxidation state of
the xanthophyll cycle; DW, dry weight; F0, initial Chl fluorescence; Fm, maximum
Chl fluorescence; Fv/F0, variable-to-initial Chl fluorescence; Fv/Fm, variable-to-
maximum Chl fluorescence; FW, fresh weight; GR, glutathione reductase; gs ,
stomatal conductance; GS, glutamine synthetase; HL, high light; HN, high nitro-
gen; LL, low light; LN, low nitrogen; MDA, malondialdehyde; N, nitrogen; P, fraction
of the absorbed PAR used in photochemistry; PAR, photosynthetically active radia-
tion; PE, fraction of the absorbed PAR neither used in photochemistry nor dissipated
thermally; PN, net carbon assimilation rate; PS, photosystem; ROS, reactive oxy-
gen species; SOD, superoxide dismutase; V, violaxanthin; Z, zeaxanthin; -GH,
-glutamyl-hydroxamate.
∗
Corresponding author. Tel.: +55 31 3899 1291; fax: +55 31 3899 2580.
E-mail address: fdamatta@ufv.br (F.M. DaMatta).
1
Present address: Departamento de Botânica, Universidade Federal de Pernam-
buco, 50670-901 Recife, PE, Brazil.
Introduction
Leaves cannot utilize all of the photosynthetically active
radiation (PAR) absorbed during exposure to full sunlight for photo-
synthesis, resulting in what is often described as excess excitation
energy (Müller et al., 2001). To protect against this excess energy,
plants can avoid light absorption, e.g., through low chlorophyll (Chl)
contents and steep inclinations to and large reflectance of incident
radiation (Adams et al., 2004). Nonetheless, if excess excitation
energy arises, it can potentially lead to the production of reactive
oxygen species (ROS) that can oxidize pigments and proteins of
the photosynthetic machinery, particularly the D1 protein subunit
of the photosystem (PS) II reaction center (Asada, 1999). To avoid
photooxidative damage, plants possess complex photoprotective
mechanisms. One key mechanism involves the dissipation of excess
absorbed light as heat in the antenna pigment complexes of PSII.
This process is related to the xanthophyll cycle, in which violax-
anthin (V) is de-epoxidated to antheraxanthin (A) and zeaxanthin
(Z). The de-epoxidation state (DEPS) of this cycle safely dissipates
0176-1617/$ – see front matter © 2010 Elsevier GmbH. All rights reserved.
doi:10.1016/j.jplph.2010.03.001